Asthenopus crenulatus Molineri, Cruz & Emmerich, 2011 (original designation).

Hubbardipes crenulatus (Molineri, Cruz & Emmerich) comb. n.

Eight autapomorphies define the genus Hubbardipes in our cladistic analysis (Appendix 2), some of them are: 1) male foretibia very wide apically (Fig. 20D), first tarsal segment partially fused with second tarsal segment (Fig. 7B); 2) thumb (inner basal projection) of penes reduced, indistinguishable (Fig. 7A); 3) apex of penis lobe wider than base and with many marginal spines (Fig. 7A); and 4) nymphal mandibular tusk with smooth to slightly crenulated inner margin, without submedian tubercles (Figs 5B, D). Additionally, the following combination of characters is useful to distinguish Hubbardipes from other genera in Polymitarcyidae: 1) ratio length male FW/foreleg = 1.1; 2) male foreleg with large and apically widening tibia, tarsal segment 1 small and partially fused to tarsal segment 2, tarsal claws slender slightly wider at apex; 3) pronotum width/length ratio: 1.5 (male), 2.2 (female); 4) marginal intercalary veins present on the entire margin of fore and hind wings, generally shorter than distance between longitudinal veins in male, but longer and anastomosed in female; 5) in both sexes FW with 3–4 crossveins between R and M, basally to R fork; 6) basal relation of FW veins IMP-MP₁ variable (IMP joined to MP₁, or basally free), MP₂ curved toward CuA and fused to CuA and MP₁ by cross veins (forming a characteristic oblique Y); 7) median remnant of styliger plate subquadrate and small, pedestals short also subquadrate and relatively small; 8) forceps slender, ratio length/basal width = 8.5–9.0 (Fig. 7A); 9) penes relatively thin, with many spines near the apex on outer edge, slightly curved inward (Fig. 7A); 10) female abdominal sternum VIII with anteromedian paired sockets on a protruding subcircular structure; 11) eggs with relatively large polar caps (almost as wide as egg, ratio width egg/cap 1.2–1.4), each cap formed by 4–7 threads, chorion completely covered by large disk-like plates and smaller irregular plates (Figs 7D–E); 12) nymphal head with a small median projection on the frons (arrow in Fig. 5C); 13) nymphs with long robust tusks, without inner tubercles, with 2 or 3-pointed apex (right and left mandible respectively) (Figs 5A–B, D); 14) nymphal foretarsal claw with single row of about 14 denticles, denticles are small basally, and larger medially, 3 denticles are closer together near the apex, and the last one is much smaller (Fig. 6D); 15) nymphal dorsal apex of hind femur with ca. 20 stout spines (Fig. 6E).

Male imago. Length (mm): body, 7.0–7.8; FW, 7.5–8.6; HW, 3.1–3.7; foreleg, 6.2–6.9; cerci, 21.6. Antennae: scape slightly longer than pedicel; flagellum bristle-like. Thorax. Pronotum width/length: 1.5–2.5. Legs. Forelegs relatively long, ratio length FW/foreleg = 1.1; tarsal segment 1 fused to tarsal segment 2 (Fig. 20D), longest segment is tibia, ratio length tarsal segment 2/tibia = 0.8; tarsal segments long decreasing in length from 2>3>4>5 (Fig. 7B); claws different in size, one long the other short, not strongly widened distally (Fig. 7C). Wings. FW with 14 marginal intercalaries along hind margin, also present along entire hind margin of HW; these intercalaries present relatively numerous connections with other cross and longitudinal veins but they are not very anastomosed; 3–4 crossveins between R and M sectors basally to R fork; Rs stem length/Rs from fork to margin = 0.24; ratio MA from fork to margin/stem length = 9–12; IMP fused basally to MP₁; MP₂ fused to CuA. Genitalia (Fig. 7A): median remnant of styliger plate present, and with pedestals short and subquadrate with inner apical corner slightly protruding distally; forceps relatively long and slender, ratio length/basal-width = 9. Terminal filament reduced, cerci long (ratio length FW/cercus = 0.35).

Female adult. Length (mm): body, 10.2–10.8; FW, 11.1; HW, 4.3. Thorax. Pronotum width/length = 1.5–2.3. Wings with more crossveins and intercalaries than male. Abdominal sternum VIII with paired anteromedian sockets on an oval and ventrally protruding structure, sockets small, shallow and contiguous. Terminal filament reduced, shorter than tergum VIII, with few thin annuli; cercus 0.5–0.6 times the length of abdomen.

Eggs (Fig. 7D–E). Length, 221–266 μ; width, 143–152 μ. Oval (ratio maximum length / maximum width = 1.5–1.7), with two medium sized polar caps on apices (ratio maximum with of egg/maximum width of coiled polar cap = 1.2–1.4), each cap formed by 4–7 very long filaments. Chorionic surface completely covered by plates: large disk-like structures frequently 3-partited with a fine microsculpture forming a dashed pattern, and many small (and irregular in shape) plates covering completely the spaces between the large plates.

Nymphs (Fig. 4A). Length (mm): body, 11.0–14.5; cerci, 4.5–5.5; terminal filament, 6.0. Head (Figs 5A, C) subquadrate in dorsal view, smooth (without pilose area), antennae 1.7–2.0 times length of head. Occipital region well developed, convex (Fig. 5A). Head capsule dorsally projected at bases of antennae. Frontal ridge marked only by a dense transversal row of setae; frons acutely projected medially (Fig. 5C); clypeus and labrum membranous and small, labrum densely covered with long setae on dorsum. Mandibular tusks (Figs 5A, B, D) relatively long and slender, similar in length to head capsule, dorso-ventrally flattened, left tusk apically with 3 tubercles (the median is reduced in length), right tusk with 2 tubercles; dorsal surface of tusks wide, with crenulated inner margin bearing long setae; outer margin with a small dorsal tubercle near base (“b” in Figs 5A–B) and densely covered with stout setae along entire margin; the small basal tubercle (“b”) forms an additional articulation between mandible and head capsule (“a” in Fig. 5A). Body of mandible: molae and canines present but small, margin between them sharp-edged (acutely protruding in right mandible, Fig. 5D); with basal U-row of long filtering setae in both mandibles. Thorax. Pronotum with short anterior ring (collar), 1/3 the length of posterior ring (length taken at the medio-longitudinal line), anterolateral corners projected, spine-like. Legs (Fig. 6). Leg I (Fig. 6A): femora very wide, well developed, with a double ventro-basal row of long filtering setae; tibio-tarsus (fused) with 3 rows of filtering setae (2 on anterior face and 1 on inner margin), roundly projected apically; tarsal claw long and slender with a row of marginal denticles (Fig. 6D). Leg II (Fig. 6B): smaller, with thinner femora, with scattered long setae, mostly basally and along hind margin; tibia and tarsi with row of long setae on outer (dorsal) margin, ventrally with many stout spines on apical half, with a distal brush of thick setae; tarsal claw weaker, without denticles. Leg III (Fig. 6C): as leg II except larger and with anterior margin of femur densely covered with thick setae, femur distally with a group of acute stout spines (Fig. 6E), tibia without distal brush. Coxae I and II directed ventrally, coxae III directed laterally. Abdomen. Sternite I stronger and partially fused with metasternum. Gill I reduced in size, dark gray, double, both portions of a similar length, but the dorsal is wider (arrow in Fig. 5E). Gills II–VII well developed, ventral portion smaller than dorsal portion. Tergum X with short and blunt, poorly developed posterolateral spine. Cerci slightly shorter than terminal filament, with long setae at joinings.

Figure 4. 

Asthenopodinae nymphs: A Hubbardipes crenulatus B Asthenopodes chumuco C Asthenopus magnus D Asthenopus angelae E Asthenopus curtus F Asthenopodes chumuco.

Figure 5. 

Hubbardipes crenulatus, nymph. A head, d.v. B left mandible, d.v. C head, frontal view (arrow indicates frontal anteromedian projection) D right mandible, d.v. E wingbuds and gill I F frontal view of thorax (head removed).

Figure 6. 

Hubbardipes crenulatus, nymph. A foreleg, d.v. (functionally frontal) B middle leg, d.v. C hind leg, v.v. (functionally dorsal) D foretarsal claw, detail E detail of apical spines on hind femur.

Figure 7. 

Hubbardipes crenulatus, male imago and egg: A male genitalia, v.v. B male foreleg C detail of foretarsal claw D egg, general view E detail of chorion (LD = large disks).

Hubbardipes from “Hubbard” and “pes”, Latin, masculine, meaning “foot”. We dedicate the genus to Mike Hubbard, mayfly specialist, who devoted many of his works to the Polymitarcyids.

Amazonas subregion (Amazonas river in Colombia and Brazil).

Hubbardipes was recovered as sister to a larger clade containing Priasthenopus, Asthenopus, Povilla and Languidipes (see synapomorphies in Appendix 2), not related with Asthenopodes, as previously thought (Molineri et al. 2011). Hubbardipes shows many differences in the adult stage and, more markedly in the nymph and egg here described for the first time. Male genitalia is unique in form and structure (Molineri et al. 2011), nymphal tusks and microsculpture on disk-like structures of the egg are exceptional as well.

Listed in Molineri et al. (2011), from Brazil (Amazonas, Presidente Figueiredo) and Colombia (Amazonas, Leticia, Reserva Natural Palmarí). Additional material: 2 male subimagos and 3 nymphs from BRAZIL: Amazonas, Tefé, São João do Catuaí, Igarapé Jutaí (A07), S 3°41'52.8" - W 64°9'18", 12.ix.2003, luz UV1, 067 FCM (CZNC); 7 nymphs from COLOMBIA, Dpto. Amazonas, rio Yavari junction Orejon, S 4°7'12" - W 69°55'43", E. Domínguez & N. Torres col. (IBN).

Hubbardipes crenulatus (Molineri et al., 2011) comb. n. is known from adults of both sexes, eggs and nymphs, and for the moment it is the only known species in the genus. The characters useful to distinguish it from other Asthenopodinae are listed in the generic diagnosis.

Nymphs (Fig. 4A). Length (mm): body, 11.0–14.5; cerci, 4.5–5.5; terminal filament, 6.0. General coloration whitish light brown. Head with a gray band between lateral ocelli and fine netting pattern on occiput (Fig. 5A). Antennae: scape bare, long and slender, pedicel shorter with many dorsal setae, flagellum bare with numerous annuli increasing in length distally. Thorax. Pronotum shaded black on anterior ring and more slightly shaded witn brownish-gray on posterior ring except on a pair of submedian longitudinal pale lines. Meso- and metanotum shaded widely with gray, with dark gray wingbuds, developing veins paler (Fig. 5E). Legs (Fig. 6). All coxae and trochanters shaded with gray, femur I shaded with gray, remaining segments and legs II and III whitish-yellow; foretarsal claw with ca. 20 denticles increasing in size distally. Abdomen. Terga more or less uniformly shaded brownish-gray, except on pale transverse dashes laterally, and pale subcircular submedian marks; tergum X with two submedian pale bands; sterna paler than terga, shaded with gray very slightly, somewhat darker on sterna IX–X. Gill I dark gray, gills II–VII brownish gray, ventral portion paler than dorsal portion. Caudal filaments whitish.

Amazonas river, from Leticia (Colombia) to Manaus (Brazil).

Hubbardipes crenulatus (Molineri et al., 2011) comb. n. was recently described from male and female adults in the genus Asthenopus, with the knowledge of the nymphs and based on the results of the phylogenetic analysis, it became evident that this species pertain to a distinct group, that we propose here as a new genus.

Asthenopus gilliesi Dominguez, 1988 (original designation).

Priasthenopus gilliesi (Dominguez), 1988 comb. n.

Priasthenopus gen. n. presents seven autapomorphies in our cladistic analysis (Appendix 2), six are variations in continuous characters (e.g., stoutter penis lobe and larger thumb) and the seventh is the presence of a short closed cleft between the aforementioned structures. This genus can be distinguished from the other by the following combination of characters: 1) ratio length male FW/foreleg = 1.6–2.0; 2) first tarsal segment subquadrate not fused with tibia (Fig. 20E); 3) pronotum width/length ratio: 2.1–2.5 (male), 2.6 (female); 4) 5–10 marginal intercalary veins present on the margin of FW (Figs 8F, H), about as long as the distance between longitudinal veins in male, slightly longer in female, HW without marginal intercalaries; 5) male FW with 1 cross vein between Rs and MA basal to Rs fork (2 in female); 6) vein MP₁ basally free (types from Uruguay and specimens from Colombia, Fig. 8D) or tending to fuse, although not completely, with MP₂ (in specimens from Bolivia, Fig. 8E), IMP basally free; 7) median remnant of styliger plate rectangular thin and convex posteriorly, pedestals rectangular (Fig. 8A); 8) ratio total length/basal width of forceps 6.3–7.3 (Fig. 8A); 9) penes relatively short with a similar width along their length, strongly curved, without apical spine or spine very slightly marked as a subapical indentation, thumb rounded (Figs 8A–C); 10) female sternum VIII with well distinguishable anteromedian sockets (Figs 8J); 11) eggs with relatively large polar caps formed by 14–16 long threads (Fig. 8K), chorion loosely covered by medium-sized and small circular plates (Fig. 8L).

Male imago. Length (mm): body, 5.0–8.0; forewing, 5.7–8.4; hind wing, 2.6–3.9; foreleg, 3.1–4.5; cerci, 19.0–25.0. Pronotum width/length: 2.1–2.5. Wings (Figs 8D–G). FW with 5–10 marginal intercalary veins (Fig. 8F), each imv is about as long as the distance between corresponding longitudinal veins; vein MP2 basally free (Fig. 8D) or base directed towards MP1 (but not completely fused with it, Fig. 8E); IMP basally free; 1 cross vein present between Rs and MA basal to Rs fork. HW without marginal intercalaries (Fig. 8G). Legs. Forelegs about half the length of FW, middle and hind legs reduced in length and poorly sclerotized but with all the segments present and distinguishable. Genitalia (Figs 8A–C): median remnant of styliger plate rectangular, thin and convex posteriorly, pedestals rectangular and relatively small (Fig. 8A); 1-segmented forceps, ratio total length/basal width 6.3–7.3 (Fig. 8A); penes (Figs 8A–C) relatively short with a similar width along their length, strongly curved medially, and without apical spine. Cerci long and well developed, terminal filament extremely reduced as common in the family.

Female adult. Length (mm): body, 6.5; FW, 8.9; HW, 3.3; cerci, 1.5. Pronotum ring-like. Wing venation (Figs 8H–I) similar to male except FW with 2 cross vein between Rs and MA basal to Rs fork (arrows in Fig. 8H). Abdominal sternum 8 with anteromedian pair of small sockets (Fig. 8J). Cerci short, about 0.2 the length of FW.

Eggs (Figs 8K–L). Length, 240–275 µ; width, 150–165 µ. Two large polar caps (maximum width, 140–185 µ), formed by 14–16 very long coiled threads. The caps are as wide as or wider than the egg. Chorionic surface with medium sized and small subcircular disks.

Figure 8. 

Priasthenopus gilliesi, adults and egg. Male imago: A genitalia, v.v. (Colombian specimen) B penes (Bolivian specimen) C penes (Uruguayan paratype) D–E FW detail of base (arrows indicate veins IMP and MP2) F detail of FW, posterior margin G hind wing. Female subimago: H forewing I hind wing J sternum VIII and detail of sockets (s). Egg (SEM): K general view L detail of cap and chorion, LD = large disks, SD = small disks.

Arbitrary combination of letters.

Treating this sole species in a new genus, distinct from Asthenopus is justified by its phylogenetic position (sister to the clade Povilla-Asthenopus). The other possibility to fit taxonomy to phylogeny would be to synonymize the entire clade (including Povilla and Languidipes besides gilliesi) in Asthenopus. This is the scheme apparently presented by Kluge (2004) but this seems inadequate to us because of relatively large morphological gaps between the groups (including characters from eggs, nymphs and adults of both sexes). Furthermore, we tested P. gilliesi position considering the hypotethical situation that its nymphs (still unknown) be identical to Asthenopus s.s., since male genitalia of both groups are similar. Even so, there were no changes in the resulting tree. The description of female adults and eggs is an original contribution of this work that gives additional diagnostic characters to genus level.

Paratype male (IFML TEPH095, slide 041) from URUGUAY, Artigas, San Gregorio, orillas río Uruguay, 29.xi.1959, light trap, C.S. Carbonell col.; 5 male imagos (slide IBN141CM) and 1 male and 1 female (slide IBN471CM) subimagos (IBN) from COLOMBIA, Amazonas, P. N. Amacayacu, río Amacayacu, 93 m, S 3°48'28" − W 70°15'21", 3.ii.1999, light trap 18−20 h PM, E. Domínguez, M.C. Zúñiga & C. Molineri cols.; 5 male imagos and 1 female subimago (IBN) from BOLIVIA, Santa Cruz, near Once Por Ciento, río Blanco, 250 m, S 15°21'39.7" − W 63°17'28.8", 14.vi.2000, light trap, E. Domínguez col.; 1 female adult (CZNC) from BRAZIL, Rio N. Aripuanã, Rio Juma, Ig. Campineiro Gde., 8−9.ix.2004, Pennsylvãnia light trap; and 2 male imagos (CZNC) from Amazonas, Barcelos, rio Demene, ´boca´barco, 8−9.viii.2009, Pennsylvania light trap.

Priasthenopus gilliesi is known from adults of both sexes and eggs, and is the only species known in the genus. The characters useful to distinguish it from other Asthenopodinae are listed in the generic diagnosis.

Male imago. See generic section above and original description in Domínguez (1988).

Female subimago. Length (mm): body, 6.5; FW, 8.9; HW, 3.3; cerci, 1.5. General coloration yellowish white with gray markings. Head cream extensively shaded gray dorsally, the shading is uniform anteriorly but in the form of a fine netted pattern posteriorly to lateral ocelli, occiput with a pair of submedian pale anterior spots and a pair of submedian dark posterior spots; venter of head whitish. Antennae yellowish white shaded with gray on scape and pedicel. Thorax cream. Anterior ring of prothorax very thin, less than 1/4 the dorsal length of posterior ring; ratio width/length: 2.6; pronotum shaded blackish on median area except pale medial line, presternum paler, shaded gray before coxa. Mesonotum shaded very diffusely with gray, darker on longitudinal carinae and between posteroscutal protuberances; mesosternum and pleurae paler, shaded gray on anterior corner of katepisternum. Metanotum shaded gray on posterior half, except on a pale median triangular mark, shaded darker posteriorly to this pale mark; metasternum whitish. Legs whitish except coxae yellowish shaded gray and apex of hind trochanter pointed and yellowish orange. Wings (Figs 8H–I). Membrane of fore and hind wings slightly shaded with light brown, veins translucent shaded with light brown, markedly on larger veins (C, Sc, Rs); marginal intercalaries relatively long (Fig. 8H), MP₁ fused with MP₂ at base; 2 crossveins present between M and R stems basally to Rs fork. Abdomen whitish, shaded with gray dorsally and darkening posteriorly, shading on terga interrupted on medial paler line, this line is wide on terga 1–4, narrows posteriorly on 5–7, thin on 8, and widens posteriorly on 9–10; sterna whitish translucent except gill sclerites yellowish white; sternum 8 with anteromedian pair of relatively large sockets (Fig. 8J). Cerci whitish, about 0.2 the length of FW.

Eggs. See generic description.

This species presents a wide geographic range that spans from the Amazon River in the North to the Uruguay River in the South, also extending towards the West in Bolivian Chiquitania.

Priasthenopus gilliesi male imagos were adequately described by Domínguez (1988), females and eggs are described here for the first time. There are no morphological differences between the male imagos examined from the different localities, except for the penes of the Colombian males are slightly stouter, and those from the Bolivian males are slightly slender than the penes of the Uruguayan types. In the forewings, vein MP₁ is basally free except on Bolivian males where this vein tends to fuse with MP₂, although not completely.

None.

In the forewings of Priasthenopus gilliesi, vein MP₁ is basally free except on Bolivian males where this vein tends to fuse with MP₂, although not completely. This last arrangement of the MP sector is also present in Campsurus paraquarius Navas (1920) and, also coincident, are the length and arrangement of the two imv (intercalary marginal veinlets) figured by Navás, the relatively short ICu veins and the vein AA basally curved to CuP; also the small size of the male described by Navás coincide with Priasthenopus size range. Navás described the color of legs without saying that middle and hind legs are reduced (as Campsurus), so probably they were present and complete as in all Asthenopodinae. Finally Navás stated that the pronotum is wider than long (transverse), feature also present in Priasthenopus and related genera (Asthenopus, Povilla) but not in Campsurus males. Other species of Neotropical Asthenopodinae show shorter intercalary marginal veins (Asthenopus s.s.) or longer ICu veins (Asthenopodes). Lestage (1923) noted the similiraty of Navás species with the genus Asthenopus and Kluge (2004) treated this species in Asthenopus because of the arrangement of veins in the Cu sector of FW. We here coincide with these authors and because of the bad original description by Navás, and the apparent loss of type material, we treat the name paraquarius (Navás 1920) as a NOMEN NUDUM.

Palingenia albicans Pictet, original designation (= Asthenopodes picteti Hubbard)

Asthenopodes picteti Hubbard, A. traverae sp. n., A. chumuco sp. n.

Seven autapomorphies define the genus Asthenopodes in our cladistic analysis (Appendix 2), some of them include: 1) Male foretarsal segment 1 fused with tibia (Figs 20A–C); 2) apex of male foretarsal claw strongly expanded (apex 3 times wider than stalk, Fig. 13E–F); and 3) pedestal relatively large, elongated, narrow at the base (Fig. 12). The following combination of characters representing the entire range of variation of the three included species, is useful to distinguish Asthenopodes from other genera in Polymitarcyidae: 1) ratio length male FW/foreleg = 1.0–1.6; 2) tarsal segment 1 indistinct (fused to tibia), tibia distally subequal in width to base of tarsal segment 2 (Figs 20A–C); 3) pronotum width/length ratio: 1.2–1.9 (male), 1.5–2.3 (female); 4) in both sexes FW marginal intercalary veins relatively long and anastomosed (from 9 to 22 in male FW, Fig. 11); 5) in both sexes FW with 3–6 (most commonly 4, but variable depending in size of specimen) crossveins between R and M, basally to R fork; 6) FW vein IMP slightly longer than MP₁, both frequently free at base but may be joined to one another and to MP₂ by crossveins (Fig. 11); 7) median remnant of styliger plate present in A. chumuco (subrectangular as other Asthenopodinae) but medially very short and with a strongly marked lateral lobe in the sister pair A. picteti-A. traverae; pedestals relatively large and thinner at the base; 8) forceps relatively slender, ratio length/basal-width = 4.7–9.5; 9) penes variable in form but curved medially rather than ventrally, basal thumb well separated from penial lobe, penial lobe apically acute (Fig. 12); 10) female abdominal sternum VIII with anteromedian paired sockets reduced in size (Fig. 13D); 11) eggs (Fig. 13A–C) with relatively small polar caps subequal to much thinner than egg, formed by 5–16 threads, chorion covered with medium-sized circular plates, surrounded by many smaller plates; 12) nymphal head dorsally strongly convex on occiput, frons not projecting medially (Figs 9A–B); 13) nymphs with very short and robust tusks (Figs 9C–H), without large submedian inner tubercle, with 2 or 3-pointed apex (asymmetric); 14) nymphal foretarsal claw with 2 rows of marginal denticles, a long row of 15 denticle and one shorter row of 12 denticles (Fig. 10B); 15) apex of femur dorsally with ca. 30 strong and rounded spines (Fig. 10D).

Male imago. Length (mm): body, 7.3–13.5; FW, 7.0–14.5; HW, 3.7–7.3; foreleg, 5.0–14; cerci, 20.0–38.5. Antennae: scape subequal to pedicel; flagellum bristle-like. Thorax. Pronotum width/length: 1.2–1.9. Legs. Forelegs relatively long, ratio length FW/foreleg = 1.0–1.6; tarsal segment 1 fused to tibiae (Fig. 20A–C); longest segment is tarsal segment 2 or tibia (variable), ratio length tarsal segment 2/tibia = 0.6–1.5; tarsal segments long decreasing in length from 2>3>4>5; claws different in size, one long the other short, strongly widened distally (Fig. 13E–F). Wings (Fig. 11). FW with 9–22 marginal intercalaries along hind margin, also present along entire hind margin of HW; these intercalaries present numerous connections with other cross and longitudinal veins (i.e., very anastomosed); 3–6 crossveins between R and M sectors basally to R fork; Rs stem length/Rs from fork to margin = 0.2–0.4; ratio MA from fork to margin/stem length = 9–15; IMP fused basally to MP₁ or free; MP₂ fused to IMP. Genitalia (Fig. 12): median remnant of styliger plate with posterolateral corners roundly projecting, pedestal long to very long; forceps relatively long and slender, ratio length/basal-width = 4.7–6.7. Terminal filament reduced, cerci long (ratio length FW/cercus = 0.32–0.44).

Female length (mm): body, 7.2–19.0; FW, 12.2–22.5; HW, 5.3–11.5; cerci 1.2–4.0. Thorax. Pronotum width/length = 1.5–2.3. Wings with crossveins and intercalaries more numerous than in male. Abdominal sternum VIII (Fig. 13D) with reduced paired anteromedian sockets, sockets small, shallow and contiguous located at the base of a blunt subquadrate projection. Terminal filament reduced, shorter than tergum VIII, with few thin annuli; cercus very short, 0.1–0.2 times the length of FW.

Eggs (Fig. 13A–C). Length, 221–355 μ; width, 143–240 μ. Oval (ratio maximum length / maximum width = 1.3–1.7), with two relatively small polar caps on apices (ratio maximum with of egg/maximum width of uncoiled polar cap = 1.2–3.1), each formed by 5–16 long coiled filaments. Chorionic surface with fine granulated aspect or regularly spaced plates.

Nymphs, nearly mature (Fig. 4B, F). Length (mm): body, 7.8; cerci, 2.0–2.3; terminal filament, 3.1. Head subquadrate in dorsal view, smooth (without pilose areas), antennae subequal in length to head. Occipital region well developed, convex (Figs 9A–B). Head capsule dorsally projecting at bases of antennae. Frontal ridge marked only by a dense transversal row of setae; frons not projecting medially; clypeus and labrum membranous and small, labrum densely covered with long setae on dorsum. Mandibular tusks very short and robust, the part visible in dorsal view ca. 1/3 the length of head capsule; left tusk (Fig. 9E–H) apically with 3 teeth, the median reduced in length, and the innermost is strongly widened appearing as a ridge with two points (Fig. 9F); right tusk (Fig. 9C–D, G–H) with 2 distal teeth; inner surface with a small tubercles located distally (in relation to other Asthenopodinae), dorsal surface with a small tubercle (“b” in Fig. 9G) that forms an additional articulation between mandible and head capsule; tusk densely covered with long setae, except at apex. Body of mandible: molae strongly protruded medially, canines present but small, margin between them sharp-edged (acutely protruding in right mandible); with basal U-row of long filtering setae in both mandibles. Thorax. Pronotum with anterior ring (collar) subequal in length to posterior ring (length taken at the medio-longitudinal line), anterolateral corners projecting, spine-like, posterior ring with dense patches of short setae medially (Figs 4F, 9B). Legs (Fig. 10A–D). Leg I (Fig. 10A): femora robust, relatively slender, with a U-shaped ventro-basal row of long filtering setae, distal points of the U almost touching each other; tibio-tarsus (fused) with 2 U-rows of filtering setae: 1 on anterior face (each branch well separated in the base) and 1 on inner margin, with the branches near each other, apex of tibio-tarsus relatively pointed; tarsal claw with 2 rows of marginal denticles (Fig. 10B). Leg II (Fig. 10C): smaller, with thinner femora, with scattered long setae basally and a row of long and short setae along outer margin; tibia and tarsi with row of long setae on outer (dorsal) margin, ventrally with many stout spines on apical half, anterior face of tibia with a distal row of thick setae (at base of tarsus) and with a crown of thick setae at apex; tarsal claw weaker, without denticles. Leg III (Fig. 10D): intermediate in size, outer margin of femur with row of short setae, longer at apex, distal corner of femur densely covered with thick, blunt setae; inner margin of femur, tibia and tarsus densely covered with short setae; margin between tibia and tarsus with row of thick setae; outer margin of tibia and tarsus with row of long pectinated setae. Coxae I and II directed ventrally, coxa III directed postero-laterally. Abdomen. Sternite I longer than the others and partially fused with metasternum. Gill I reduced in size, whitish, single and lanceolate. Gills II–VII well developed, ventral portion smaller than dorsal portion. Tergum X without posterolateral spine (Fig. 10E). Cerci slightly shorter than terminal filament, with long setae at joinings, basal 1/4 with thick blunt setae ventraly (Fig. 10F).

Figure 9. 

Asthenopodes nymph. A. chumuco: A head, dorsal view B head and pronotum, d.v. C–D right mandible and detail of apex, v.v. (arrow = subdistal tubercle) E–F left mandible and detail of apex, v.v. G mandibles, d.v. (b = dorsal tubercle) H same, v.v. A, G, H stereomicroscope photographs B–F SEM.

Figure 10. 

Asthenopodes nymph. A. chumuco: A foreleg, v.v. B detail of foretarsal claw C middle leg, d.v. D hind leg, d.v. with detail of femoral apex E abdominal sterna IX–X (arrow indicate absence of spine on paraproct) F detail of cercus. All figs from SEM except A from stereomicroscope.

Figure 11. 

Asthenopodes wings, steromicroscope photographs. Males to the left, females to the right: A–D A. picteti E–H A. traverae I–L A. chumuco. Not to scale.

Figure 12. 

Asthenopodes, male genitalia: general view to the left, detail of penes to the right. A–B A. picteti C–D A. traverae E–F A. chumuco. Scale bar = 200 µ in E 100 µ in F. Abbreviations: mp = median plate of stryliger; pe = pedestal; s9 = ninth sternum; see Appendix 2 for explanation of measures (letters a to g, and z).

Figure 13. 

Asthenopodes, egg and adults. Eggs: A A. picteti B A. traverae C A. chumuco. Female imago: D A. traverae, abdominal sternum VIII, and detail of keel (k) and sockets (s). Male imago foretarsal claw: E A. chumuco F A. traverae.

Amazonas and Parana subregions (Argentina, Brazil, Colombia, Guyana, Uruguay).

Asthenopodes and Asthenopus have been treated as synonyms (Hubbard and Domínguez 1988) after the discovery of Priasthenopus gilliesi (Domínguez, 1988), that somewhat blurred the distinction between both genera. Additionally, Hubbard and Domínguez (1988) based their synonymic proposal in the fact that all known nymphs of Asthenopodinae from South America where indistinguishable and could be classified in a single genus. As the knowledge of this group had largely improved in last years we are proposing here a new rearrengement of supraspecific taxa.

The characteristics traditionally associated with Asthenopodes (summarized in Domínguez 1988) are: 1) ratio foreleg/forewing of male: 7/8; 2) male foretarsus 3.5 times longer than foretibia; 3) foretarsal segment 2 very long (almost as long as tarsal segments 3 and 4 combined, and 1.5 times the length of tibia); 4) Rs fork base to margin: 2.5/10; 5) cubital intercalaries parallel, ICu₂ arising basally from ICu₁, or basally free but connected to ICu₁ and CuP by cross veins, 6) ICu₂ ending at hind margin, 7) long marginal intercalary veins present; 8) forceps ratio width/length: 1/10; 9) penes thin from the base; 10) MA fork base to margin 7/100; 11) IMP–MP₁ fused; 12) MP₂-IMP similar in length, fused; 13) foretarsal claws of male greatly expanded apically. The discovery of P. gilliesi (Domínguez) put in doubt the value of characters 5, 7, 8, 9 and 10, for generic diagnoses, because this species showed some intermediary states between Asthenopodes and Asthenopus (Domínguez 1988, Hubbard and Domínguez 1988). In our phylogenetic analysis the intermediate position of P. gilliesi is mantained, but it is clearly located outside both genera.

The revalidation of the genus Asthenopodes Ulmer is based not only in the clade that its type species (Asthenopodes picteti) forms with other two new species, but also on the fact that the nymph shows characters considered important at the generic level in the family, mainly the shape of nymphal mandibular tusks, legs and gill I.

Male imagos

Female and eggs

Type material was not studied; it consists of the holotype male imago, damaged, with many parts missing including the genitalia. It is a pinned specimen deposited at Naturistorisches Museum Wien, Hubbard and Domínguez (1988) presented a figure of the holotype forewing.

Additional material. 3 male subimagos from ARGENTINA, Misiones, Parque Provincial Urugua-í, Arroyo Yacutinga, S 25°44'51" − W 54°03'37", 355 m, 30.xi.2001, Domínguez et al. cols.; 8 female and 2 male imagos same data except Arroyo Uruzú, S 25°51'29" − W 54°10'10", 322 m, 25.xi−2.xii.2001; 1 female and 1 male imagos (slides IBN3–93 and 3–96) from URUGUAY, Maldonado, Arroyo de la Quinta, 4.i.1984, M. T. Gillies col. All the material deposited in IBN.

Asthenopodes picteti, type species of the genus Asthenopodes (Ulmer 1924, Traver 1956, Hubbard 1975) presents 7 autapomorphies (Appendix 2) including: thinner male foretibia and slender penes. This species can be distinguished from the other species of the genus by the following combination of characters: 1) general coloration yellowish white (male), darker in female (yellowish light brown); 2) male FW 11.4–11.9 mm, female FW 14.5–19.0 mm, membrane whitish hyaline tinged with yellowish near hind margin; 3) foreleg length 0.9 times the length of FW; 4) pronotum width/length ratio: 1.6–1.9 (male), 1.5–2.3 (female); 5) 14–17 marginal intercalary veins present on the entire margin of FW (Fig. 11A) and HW (Fig. 11B), miv generally longer than distance between longitudinal veins; 6) male FW with 4 to 6 cross veins between Rs and MA basal to Rs fork; 7) ratio total length/basal width of forceps 6.7 (Fig. 12A); 8) penes long and slender (Fig. 12A–B), male median remnant of styliger plate posterolaterally expanded forming a pair of rounded projections (“Ll” in Fig. 12B); 9) female sternum VIII, anteromedian sockets present but concolorous with sternum, not evident; 10) egg polar caps only slightly thinner than maximum width of the egg, formed by 6 filaments; medium-sized and small chorionic disks-like structures present (Fig. 13A).

Male imago. Length (mm): body, 9.3–11.2; FW, 11.4–11.9; HW, 5.8–6.2; foreleg, 10.0–10.3; cerci, 30.1. Described in Traver (1956) and Domínguez (1988a). Additions to these descriptions follow: Prothorax width/length: 1.6–1.9. Wings. Hind margin of FW (Fig. 11A) with 14–17 marginal intercalary veins, relatively long; 4–6 cross veins between R and M, basal to R stem; IMP basally free or fused to MP₁ by a cross veins, MP₂ fused to IMP. HW as in Fig. 11B. Genitalia (Fig. 12A–B): forceps length/width ratio: 6.7; median remnant of styliger plate with posterolateral corners roundly projecting.

Female adult. Length (mm): body, 11.5–12.6; FW, 14.5–19.0; HW, 6.3–9.4. General coloration yellowish light brown. Head dorsally blackish except on median zone, paler; venter of head yellowish white. Antennae light brown, shaded gray on scape. Thorax yellowish brown with blackish membranes, shaded with brownish gray on pronotum and with black on posteromedian marks on meso- and metanotum. Pronotum width/length: 1.5–2.3. Legs whitish yellow shaded brownish on dorsum of leg I and on apex of femur III. Wings (Fig. 11C–D), membrane tinged with light brown, veins yellowish brown. Abdomen. Terga brownish with a pale mediolongitudinal line and paler areas on lateral margins on terga I–VII; sterna whitish yellow; female sternum VIII with anteromedian sockets, small and almost not distinguishable. Terminal filament whitish, shorter than tergum VIII; cerci yellowish brown paler apically, 0.5–0.6 times the length of the abdomen (0.2 the length of FW).

Eggs (Fig. 13A). Length, 325−355 µ; width, 215−235 µ. Two large and flat polar caps (maximum width, 155−180 µ), formed by 6 very long coiled threads. Chorionic surface with a fine granulated aspect, with small disk like structures.

Uruguay, Argentina. A. picteti is here newly recorded from Argentina. The record from Guyana given by Domínguez et al. (2006, p.562) is no longer valid since this material is now considered a different species (A. chumuco see below).

Asthenopodes picteti Hubbard was only partially known from the damaged holotype male from Brazil until Traver (1956) redescribed it (at that time as Asthenopodes albicans) based on a complete male from Uruguay. Later Hubbard (1975) gave a new name to this species (A. picteti). Domínguez (1988) and Hubbard and Domínguez (1988) presented additional discussion and illustrations of the male imago, from type and non-type material (also collected in Uruguay). The description of Traver (1956) coincides with the new material studied, except for the relation of the male foretibia and forefemur. Traver reported that tibia is 1.33 times the length of femur but we found that it is 1.7 times that length.

Domínguez (1988) reported 2 female imagos collected at the same time than the males he redescribed, but did not presented a formal description of them. Both females and additional ones from Argentina (see list of material) are here shortly described and figured. It is difficult from the material available to determine if the females are in subimaginal or imaginal stage, so they are here referred as “adults”. The females are similar to the sympatric A. traverae but can be separated from this species because A. picteti females are lighter in color, hind femora only shaded black on apex, paired female sockets present but small and hard to distinguish, cerci light colored, abdominal gill sclerites smaller, FW wider with shorter marginal intercalaries joined to main veins and partially anastomosed. The eggs present wider polar caps, only slightly thinner than the egg. Males can be separated from those of A. traverae because the hind femora are only shaded black on apex and the penes are much slender.

Holotype (IBN) male and paratypes (IBN) 4 males and 22 females (slides IBN468–469CM) from ARGENTINA, Misiones, Parque Provincial Urugua-í, Arroyo Uruzú, 7−11.xii.1999, C. Molineri col.; 1 female imago (“Asthenopodes sp.” Traver det.) from URUGUAY, Artigas (Uruguay 19), 9.i.1952; and 27 female adults from URUGUAY, Artigas, Sepulturas (D-3), 18.xii.1952, Carbonell col. Paratypes from Uruguay deposited in FCE-Ep.

Non-type material: 1 female (wings on slide) from BRAZIL, Sao Paulo, Jacareí, rio Paraiba do Sul, 21.xi.1987, CG Froehlich et al. cols (deposited at MZSP).

Four autapomorphies characterize this species, all are small changes in continuous characters, except the marked shortening of the median remnant of styliger plate (at the middle, since laterally a tong-like projection is present, Fig. 12C). Asthenopodes traverae sp. n. is known in the alate and egg stages, all females are subimagos even those that apparently have laid the eggs (empty females). This species can be distinguished from the other species of the genus by the following combination of characters: 1) males yellowish white, females blackish; 2) FW length male 14.0–14.5 mm, female 15.5–22.5 mm, membrane yellowish hyaline; 3) male foreleg length 0.9–1.0 times the length of FW; 4) pronotum width/length ratio: 1.3–1.9 (male), 2.0 (female); 5) 16–22 marginal intercalary veins present on the entire margin of FW (also present in HW) generally longer than distance between longitudinal veins, fused to main veins, poorly anastomosed (Fig. 11E–H); 6) male FW with 4 to 5 cross veins between Rs and MA basal to Rs fork; 7) ratio total length/basal width of forceps 4.7–4.9 (Fig. 12C–D); 8) median remnant of styliger plate posterolaterally expanded forming a pair of rounded projections, penes large and robust, sclerotized, curved and twisted (Fig. 12C–D); 9) female sternum VIII with anteromedian sockets well developed, whitish surrounded by a large brownish area (Fig. 13D); 10) egg caps small, much thinner than maximum width of the egg, formed by 5–6 filaments; medium-sized and small chorionic plates present (Fig. 13B).

Male imago. Length (mm): body, 12.2–13.5; FW, 14.0–14.5; HW, 6.8–7.3; foreleg, 12.3–14.0; cerci, 35.5–38.5. General coloration yellowish white. Head shaded black dorsally almost entirely, with a pair of distinct submedian black marks anteriorly to median ocellus; occipital hind margin with pale median zone; head ventrally pale without markings. Antennae: scape and pedicel short, yellowish on venter of pedicel, both slightly shaded with gray; flagellum very thin, hyaline, similar in length to forefemur. Thorax. Pronotum translucent, shaded slightly with gray except on membranes separating anterior and posterior rings, darker laterally; pronotum width/length: 1.3–1.9. Meso- and metanotum yellowish white with gray markings mainly posteriorly but also on sutures. Thoracic pleurae and sterna yellowish white shaded gray only at base of coxae and wings. Legs. Forelegs: coxa dorsally whitish with a gray mark, ventrally yellowish; femur dorsally yellowish shaded gray on apical third, ventrally whitish; tibia whitish translucent shaded gray mainly on dorsum; tarsi translucent shaded slightly with gray; large claws, stalk with brown inner margin, rest whitish; articulations between femur-tibia and tibia-tarsus very sclerotized, brownish. Middle and hind legs yellowish, shaded with gray from half of femur to apex of leg. Wings (Fig. 11E–H). Membrane hyaline slightly tinged with gray on costal and subcostal sectors (membrane yellowish under transmitted light); all veins light gray, lighter toward hind margin; 5 cross veins between R stem and M sector; long marginal intercalaries on hind margin of both wings. Abdomen yellowish white widely shaded with gray, some darker marks as follows: submedian short anterior dashes and sublateral oblique stripes on terga III–VIII, lateral margins of terga VIII–IX, and median line of IX–X (thinner on X). Abdominal sterna whitish except gill sclerites yellowish white. Genitalia (Fig. 12C–D): lateral margins of sternum IX and pedestals yellowish; median remnant of styliger plate whitish with a gray strip on hind margin between the tongue-like projections (“Ll”Fig. in Fig. 12D); pedestals well separated from each other, relatively long and becoming wider distally; forceps whitish, long and wide; base of penes well developed, subquadrate, whitish; penes strongly sclerotized, orangeish basally but yellowish distally. Cerci whitish, terminal filament reduced to 7–8 thin annuli, straight, whitish.

Female subimago. Length (mm): body, 9.5–19.0; forewing, 15.5–22.5; hind wing, 7.5–11.5; cerci 3.5–4.0. General coloration dark brown shaded widely with black. Head black dorsally, yellowish white ventrally except on remnants of tusks, brownish. Antennae dark brownish except apical half of flagellum whitish. Thorax. Sclerites dark brown, membranes shaded black. Pronotum width/length: 2. Legs brownish except membranous portions, whitish. Wings (Fig. 11G–H) with yellowish brown membranes and veins, veins C and Sc darker. Abdomen. Terga dark brown completely shaded black; sterna paler, brownish laterally, yellowish medially, with two or more pairs of small pale dots; sternum VIII with two small anteromedian sockets (Fig. 13D). Caudal filaments brownish, terminal filament shorter than tergum VIII, cerci 0.3–0.4 times the length of the abdomen (0.2 the length of FW).

Eggs (Fig. 13B). Length, 305–355 µ; width, 195–225 µ. Two small polar caps (maximum width, 70–82.5 µ) formed by 5–6 long coiled threads. Chorionic surface smooth with regularly spaced chorionic plates. Chorionic plates (“disk-like” structures) relatively small and rounded, with irregular margins. Smaller plates, irregular in form scattered between the larger ones.

The species is dedicated to the great mayfly specialist Jay R Traver, who visited Uruguay and worked with Mr. C. S. Carbonell’s collections at the “Museo de la República” recognizing the females of this species as distinct from A. picteti (also unknown at that time).

Parana biogeographic subregion in Argentina, Uruguay and Brazil.

Females subimagos were collected (in Misiones Province) while swarming in compact groups at about 3 m above water in pool areas around sunset. The same behavior was reported by Traver (1956) for the Uruguayan females. Males were caught at light traps during the first hours of dark, so the male flight is unknown.

Traver (1956) described females of A. traverae as distinct from A. picteti in spite of the fact that the female of that species was also unknown at that time. Nevertheless Traver realized that they differ from A. picteti males in color and size, and left them unnamed. With the collection of new material from both species (and sexes) in Misiones (Argentina), it became evident that they constitute a new species. Asthenopodes traverae females can be distinguished from A. picteti females by their black general coloration, somewhat slender forewings, femora and cerci widely shaded with black, and enlarged abdominal gill sclerites (remnants of nymphal gill muscles insertions), for other differences see discussion under A. picteti. Very similar in aspect but much smaller are the females of A. chumuco, the allopatric sister species of A. traverae.

Holotype and 3 paratypes male imagos from Brazil, Amazonas, Barcelos, rio Demene, ´boca´barco, S 0°25'28.7" - W 62°54'20", 8−9.viii.2009, Pennsylvania. Holotype and 1 paratype in INPA, 2 paratypes in IBN.

Additional material. 3 male slides (CUIC) from British Guiana, Bartica District, Kartabo, 20.iv.1919, C.U. Expedition col. Five female imagos (1 in CZNC, 2 in IBN, IBN533CM, 2 in MUSENUV) from Colombia, Amazonas, Puerto Nariño, Loreto Yacu, S 3°44'26" − W 70°27'19", 5.ii.1999, luz 18−20 h, M. C. Zúñiga, E. Domínguez and C. Molineri cols.

Asthenopodes chumuco known from all the stages presents seven autapomorphies, all of them are changes in continuous characters (Appendix 2). This species can be distinguished from the other species of the genus by the following combination of characters: 1) general coloration yellowish white in male, dark brown in female; 2) male FW 7.0–9.0 mm, female FW 12.2–14.8 mm; 3) male, ratio FW/foreleg length 1.4–1.6; 4) pronotum width/length male 1.3–1.4, female 2.0–2.1; 5) 7–14 marginal intercalary veins present on the entire margin of FW (Fig. 11I, K), HW (Fig. 11J, L) with 3–7 marginal intercalary veins, generally longer than distance between longitudinal veins, fused to main veins, poorly to heavily anastomosed; 6) male FW (Fig. 11I) with 3–4 cross veins between Rs and MA basal to Rs fork; 7) ratio total length/basal width of forceps 7.8–9.5 (twisted CUIC slide 6.3–6.7) (Fig. 12E); 8) male median remnant of styliger plate subrectangular, slightly convex (not expanded forming a pair of rounded lateral projections), penes long and slender, acute distally (Fig. 12E–F), rectangular pedestals strongly enlarged, ½ the length of forceps; 9) female sternum VIII, with small reduced anteromedian female sockets at the base of a median keel; 10) egg caps small, much thinner than maximum width of the egg, formed by 14–16 filaments; large and small chorionic plates present (Fig. 13C).

Male imago. Length (mm): body, 7.3–8.0; forewing, 7.0–8.8; hind wing, 3.7–4.3; foreleg, 5.0–5.8; cerci, 20.0–23.0. General coloration yellowish white. Head shaded gray dorsally almost entirely, frons with black dot at base of antenna, with medial line and irregular black marks; occipital region with pale median zone; head ventrally pale without markings. Antennae: scape and pedicel short, subequal in length, whitish shaded with purplish; flagellum very thin, hyaline. Thorax. Pronotum whitish with anterior and posterior portion subequal in size, shaded gray in a transverse band between both portions, posterior portion black along hind and lateral margins; pronotum width/length: 1.2–1.3. Mesonotum yellowish white shaded with gray on posterior half of medial line, on area between posterolateral protuberances and on anterior margin of these structures. Metanotum yellowish slighlty shaded with gray medially. Thoracic pleurae and sterna yellowish white shaded gray dorsally and anteriorly to mid coxa. Legs. Forelegs: whitish completely shaded with gray; large claws, apically expanded (Fig. 13E). Middle and hind legs yellowish white, shaded with gray on coxae. Wings (Fig. 11I–J). Membrane hyaline shaded with gray on basal 1/3 of costal and subcostal sectors; all veins translucent, except costal cross veins grayish; 3–4 cross veins between R stem and M sector; long marginal intercalaries on hind margin of both wings. Abdomen whitish shaded with gray on terga, mainly on lateral margin, submedian black dot on terga III–VIII. Abdominal sterna pale. Genitalia (Fig. 12E–F): yellowish white, except for penis apically yellowish; pedestals well separated from each other, very long, ½ length of forceps; median remnant of styliger plate with slightly convex hind margin; base of penis rounded, projecting posterolaterally, lobe of penis long and sclerotized, curved ventromedially, constricted on median length, gonopore well developed. Cerci whitish; terminal filament reduced to 5–7 thin annuli, straight, whitish.

Note: Cornell male (slides, male imago). Length (mm): body missing; FW, 9.0; HW, 4.5. FW with 11 long marginal intercalary veins; 3 cross veins between R and M basad to R stem; IMP fused basally to MP₁ or free; MP₂ fused to IMP. HW with 6 long intercalary veins on hind margin. Genitalia: penes robust, twisted; median remnant of styliger plate without posterolateral projections; forceps long and slender, ratio length/basal width: 6.3–6.7.

Female imago. Length: body, 7.2 (shrunken, empty)–12.3; FW, 12.2–14.8; HW, 5.3–6.0; cerci, 1.2–1.3. General coloration dark brown. Head dorsally black except on clypeus, whitish with a pair of lateral brownish bands, ventraly much paler brownish white. Thorax brownish with blackish membranes and carinae. Pronotum with a pair of distinct black marks submedially; width/length ratio: 2. Legs pale, brownish white. Wings (Fig. 11K–L) with hyaline membrane, slightly whitish translucent; veins whitish tinged with brownish basally; 2 to 4 crossveins between M and R, basally to R stem; MP₂ joined basally to IMP; marginal intercalaries relatively numerous and long, anastomosed. Abdomen. Terga brownish slightly paler on median band and pleural folds, sterna much paler brownish white. Cerci basally brownish turning whitish distally; very reduced in size, less than 0.1 the length of FW.

Eggs. Length 210–240 µ, width 180–200 µ. Subovate, yellowish, with two small whitish polar caps (maximum width, 75–85 µ), polar caps much thinner than the egg and formed by 14–16 threads. Under SEM the larger disk-like chorionic structures are surrounded by many smaller ones, which at their time are surrounded by smooth chorion (Fig. 13C).

Nymphs. Length (mm): body, 7.8; cerci, 2.0–2.3; terminal filament, 3.1. General coloration yellowish light gray (Fig. 4B, F). Head with a black band between lateral ocelli and fine netting pattern on occiput (Fig. 4F). Antennae: scape bare, slightly longer than pedicel, pedicel with many dorsal setae, flagellum bare with numerous annuli increasing in length distally. Thorax. Pronotum shaded black on sublateral area of anterior ring and laterally on posterior ring. Meso- and metanotum shaded widely with gray, with dark gray wingbuds, developing veins paler. Legs (Fig. 10A–D). Coxae and trochanters of mid and hind legs slightly shaded with gray, remainder of legs yellowish-white; foretarsal claw with double parallel rows of 15 and 12 denticles each (Fig. 10B). Abdomen. Terga more or less uniformly shaded brownish-gray, except on pale transverse dashes laterally, thin medial line on tergum I–IX becoming wider posteriorly and pale with subcircular submedian sigilla; sterna yellowish. Gill I whitish, gills II–VII purplish gray, ventral portion paler than dorsal portion. Caudal filaments yellowish.

“Chumuco” is one of the common names applied to river cormorans in some South American countries. The penis lobe of this new species resembles the neck and head of that bird.

Brazil (Amazonas, Espírito Santo), Colombia (Amazonas), Guyana.

The male imago of this species has been known since Traver (1950) found a group of slides containing some body parts of a missing specimen. She stated that it was surely not the type species of Asthenopodes mainly because its smaller size. Domínguez et al. 2006 (p. 562) treated this specimen in Asthenopus picteti, extending the distribution of this last species to British Guiana. With the discovery of new specimens (male and female adults) we gathered more morphological information and realized that a new species must be described. One of the results is that A. picteti is no longer considered to be in British Guiana. The Colombian females described here as Asthenopodes chumuco are associated with the males, because of the egg morphology, compared with those extracted from a pharate female from São Mateus. Adults show the smallest size of the genus, further differing from the other two species in many morphological aspects. The females are similar to A. picteti in coloration of hind femur and abdomen but wing venation and sockets on sternum VIII are different, also the female cerci of A. chumuco are much less developed. The eggs are similar to those of A. traverae, because of the small polar caps, nevertheless A. chumuco presents much more threads forming each cap (between 14 and 16, but each thread is relatively very thin).

Palingenia curta Hagen, original designation.

A. curtus, A. angelae, A. magnus sp. n., A. hubbardi sp. n., A. guarani sp. n.

Five autapomorphies were recovered for Asthenopus: 1) character 1 (ratio length second foretarsal segment/foretibia) decreases from 0.645–0.652 to 0.584–0.587; 2) char. 7 (marginal connectivity = n°of connections among imv/n°of imv) decreases from 0.700–1.167 to 0.222–0.300; 3) male foretarsal segment 1 subrectangular; 4) penes with an apical spine; and 5) long and open cleft present between penial lobe and thumb. The following combination of characters is useful to distinguish Asthenopus from other genera in Polymitarcyidae: 1) ratio length male FW/foreleg = 1.4–1.8; 2) tarsal segment 1 distinct and subrectangular in form (not fused to tibia), ratio subapical width of foretibia/subbasal width of second tarsal segment 1.5–2.3 (Fig. 20F–J); 3) pronotum width/length ratio 1.7–2.4 (male, but frequently around 2.0), 2.0–3.0 (female); 4) in both sexes FW marginal intercalary veins short and not anastomosed (4–25 in number, Fig. 16); 5) in both sexes FW with 0–4 (most commonly 2, but variable depending on size of specimen) crossveins between R and M, basally to R fork; 6) FW vein IMP basally free, subequal in length to MP₂, forming a characteristic oblique “Y” with associated cross veins (arrow Fig. 16A, E); 7) median remnant of styliger plate present, pedestals well developed (Fig. 17); 8) forceps relatively stout, ratio length/basal-width = 4.7–7.0; 9) penes relatively short and stout, curved ventrally (and medially), with a thumb attached at the base, and with the apex of penial lobe distinctly shaped as a spine; 10) female abdominal sternum VIII with anteromedian paired sockets, much reduced in size, and with a keel as in Figs 18F–H; 11) eggs with relatively large polar caps formed by 3–8 (commonly 5) long coiled threads, chorion covered with disk-like structures (Fig. 18A–E); 12) head dorsally strongly convex on occiput, frons not projecting medially (Fig. 14I–J); 13) nymphs with very short and robust tusks (Fig. 14A–C, E–G), with large submedian inner tubercle, with 2 or 3-pointed apex (asymmetric); 14) nymphal foretarsal claw with a single row of denticles (Fig. 15G); 15) apex of femur 3 with a group of ca. 50 stout pointed spines (Fig. 15F).

Male imago. Length: body, 6.5–10.5; FW, 7.0–10.1; HW, 2.9–4.8; foreleg, 5.1–7.5; cerci, 22.0–37.0. Antennae: scape slightly longer than pedicel, flagellum bristle-like. Thorax. Pronotum width/length: 1.7–2.4. Legs. Forelegs subequal to shorter than body, ratio length FW/foreleg 1.4–1.8; longest segment is tibia (ratio length tarsal segment 2/tibia = 0.4–0.7); tarsal segment 1 distinct, not fused, very short (Fig. 20F–J); remaining tarsal segments relatively short subequal in length except tarsal segment 2 slightly longer; claws differing in length, one long the other short, very slightly widened distally. Wings (Fig. 16). FW with 4–25 marginal intercalaries, sometimes present in HW but not in all spaces; intercalaries not strongly anastomosed; 0–4 cross veins between MA and R, basad to R stem; R stem length/Rs from fork to margin = 0.17–0.31. Ratio MA length from fork to margin/stem length = 7–15; IMP fused basally to MP₁ or free; MP₂ fused to IMP. Genitalia (Fig. 17): median remnant of styliger plate subrectangular to subovate; pedestals subrectangular to subovate, relatively small; forceps relatively wide to very wide, ratio length/basal width 4.7–7.0; penes tubular and robust, curved ventro-medially, and with well-developed basal thumb. Terminal filament reduced, cerci long (ratio length FW/cercus = 0.25–0.36).

Female adult. Length: body, 8.0–19.5; FW, 12.0–18.5; HW, 4.6–7.8; cerci, 3.0–7.0. Thorax. Pronotum width/length = 2–3. Wings with more crossveins and intercalaries than in male. Abdominal sternum VIII with anteromedian keel (Fig. 18F–H), at each side of keel´s base a very small “socket” is present (“s” in Fig. 18F). Terminal filament reduced, shorter than tergum VIII, with few thin annuli. Cercus short, 0.25–0.50 times length of FW.

Eggs (Fig. 18A–E). Length, 210–285 µ; width, 135–163 µ. Oval (ratio maximum length / maximum width = 1.4–1.8), with two relatively large polar caps (ratio maximum with of egg/maximum width of uncoiled polar cap = 1.1–1.5), formed by 3–8 very long coiled threads. Chorionic surface with large subcircular chorionic plates, sometimes each plate is divided in 2–3 portions.

Nymphs. Length (mm): body, 9.7–15.0 mm; cerci, 4.0–7.0; terminal filament, 5.0–5.1. Head suboval in dorsal view, smooth (without pilose area); occipital region well developed, strongly convex (Figs 4C–D, 14I–J). Head capsule with a dorsal spine-like projection at bases of antennae. Antennae 1.1–1.5 times length of head (length of head taken from hind margin to the apex of clypeus); pedicel with tuft of setae on dorsum, flagellum with minute scattered setae; length (mm): scape (0.5), pedicel (0.28), flagellum (2.0). Frons with anterior margin more or less straight (arrow in Fig. 14J), with a small blunt lateral projection (“a” in Fig. 14I), without median projection. Clypeus and labrum small, membranous, with many setae on dorsum of labrum. Mandibular tusks robust, relatively stout, left tusk (Fig. 14A–C, E) with 3 apical teeth, increasing in size from the median (smallest), inner and outer; inner tooth slightly directed medially, others directed distally; right tusk (Fig. 14F–G) with 2 teeth, the inner shorter. Inner margin of both tusks with a rounded small tubercle near subapex and a larger and pointed subbasal tubercle (associated with a tuft of rigid setae), this large basal tubercle shows a small basal protuberance (giving the impression of a bifid tubercle but with one of the sides aborted); ventral surface and outer margin of tusks with small rounded protuberances on the extremely hard cuticle; dorsal surface of tusks with numerous setae and with a small basal tubercle; this small dorsal tubercle is easily seen without dissecting the mandible and gives an additional point of articulation between the mandible and the head capsule (“a” and “b” in Fig. 14I). Incisors and prostheca of both mandibles very reduced in size, molae relatively well developed. Maxillae with a small subtriangular basal membranous “gill” (membranous outgrouth). Thorax. Anterior ring of pronotum (or collar sensu Kluge 2004) short (ca. 1/4 the length of posterior ring), anteriorly projecting as spines on lateral corners; posterior ring longer, ring-like. Legs (Fig. 15A–D, F–G). Leg I (Fig. 15A–B): femora very wide, well developed, with a double ventro-basal row of long filtering setae; tibio-tarsus (fused, but fusion line distinguishable) with 3 rows of filtering setae (2 on dorsal “face” and 1 on inner margin), tarsus slightly and bluntly projecting apically (arrow in Fig. 15B); tarsal claw relatively large and stout with a row of marginal denticles (Fig. 15G). Leg II (Fig. 15C): smaller, with thinner femora, with scattered long setae, mostly basally and along hind margin; tibia and tarsi with row of long setae on outer (dorsal) margin, ventrally with many stout spines on apical half, with a distal brush of thick setae (arrow in Fig. 15C); tarsal claw relatively small, without denticles. Leg III (Fig. 15D, F): as leg II except larger and with anterior margin of femur densely covered with thick setae, and posterior margin roundly expanded at apex bearing a group of stout acute spines (Fig. 15F); tibia without distal brush. Coxae I and II directed ventrally, coxae III directed laterally. Abdomen. Gill I reduced in size, double, both portions subequal in length and width. Gills II–VII well developed, ventral portion smaller than dorsal portion; tergum X with well developed ventral spine on posterior margin (not visible dorsally, Fig. 15E). Caudal filaments short (curved in mature nymphs) with whorls of stout spines and simple setae at joinings.

Figure 14. 

Asthenopus (and Povilla) nymphs, head and mandibles. A–D left mandibles, d.v.: A A. curtus B A. angelae C A. magnus D Povilla adusta. E left mandible dorso-oclusal view, A. magnus. Right mandibles, d.v.: F A. angelae, G A. magnus H P. adusta. I–J, A. magnus head capsule, d. v., with and without mandibles.

Figure 15. 

Asthenopus nymphs. A. magnus: A foreleg, d.v. B same, v.v. (arrow indicates apical projection of tibiotarsus) C middle leg, d.v. (arrow indicates distal brush on tibia) D hind leg E abdominal sterna IX–X (arrow indicates spine on paraproct). A. angelae: F hind femur, d.v. G foretarsal claw. A–G stereomicroscope photographs F–G light microscope photographs.

Figure 16. 

Asthenopus fore (FW) and hind wings (HW) of male imago. A–B A. curtus FW & HW C–D A. magnus FW & HW E–F A. hubbardi FW & HW G–I A. guarani, FW (details) & HW J–K A. angelae (from Argentina) FW & HW. Ephoron sp.: L male HW (x.i. = extra intercalary). Stereomicroscope photographs (except L line drawing).

Figure 17. 

Asthenopus, general ventral view of male genitalia and detail of penes. A–B A. curtus C A. hubbardi D A. angelae E–F A. guarani G A. magnus. Light microscope photographs.

Figure 18. 

Asthenopus, SEM photographs. Eggs: A A. curtus (egg extracted from mature nymph), B A. angelae (egg extracted from mature nymph) CA. cf. guaraniD A. hubbardi, E A. magnus. Female abdominal sternum VIII: F A. magnus G A. curtus H A. hubbardi.

Figure 19. 

Povilla spp, SEM and light microscope photographs. P. adusta: A male forewing B male hind wing C detail of keel on female abdominal sternum 8 (b = base, k = keel, s = socket) D male genitalia (mp = median remnant of styliger plate) E–F female abdominal sternum 8 (general view) G male genitalia (median remnant of styliger plate partially broken and detached from right pedestal) H–I eggs and detail of chorion (LD = large disks). P. cf. heardi: F female abdominal sternum VIII J filaments surrounding the eggs inside female abdomen K eggs. Scale bar = 100 µ, except Figure 1 (10 µ).

Figure 20. 

Foreleg of male imago, detail of articulation between tibia and tarsus: A Asthenopodes picteti B Asthenopodes chumuco C Asthenopodes traverae D Hubbardipes crenulatus E Priasthenopus gilliesi F Asthenopus angelae G Asthenopus curtus H Asthenopus guarani I Asthenopus hubbardi J Asthenopus magnus; K Povilla adusta L Ephoron sp. Abbreviations: Ti = tibia; Ta1 = tarsal segment 1; Ta2 = tarsal segment 2.

Amazonas and Parana biogeographic subregions (Argentina, Bolivia, Brazil, Colombia, Ecuador, Peru).

The genus Asthenopus has been distinguished by means of the following characters (Domínguez 1988): 1) ratio foreleg/FW male: 3/5–4/5; 2) male foretarsus 2.5 times longer than foretibia; 3) foretarsal segment 2 similar to the others, and 2/3 the length of tibia); 4) ratio length of Rs stem/fork to margin 1/4 (or fork Rs at 2/10 from base to margin); 5) cubital intercalaries slightly diverging toward hind margin, ICu₂ and ICu₁ basally fused to CuA by cross veins; 6) ICu₂ ending at anal margin or in the tornus, 7) marginal intercalary veins absent; 8) forceps ratio width/length: 1/7; 8) penes robust on basal 2/3; 9) MA fork base to margin 10/100; 11) IMP–MP₁ not fused basally; 12) MP₂-IMP similar in length, not fused; 13) foretarsal claws of male not so expanded distally (as in Asthenopodes). Our phylogenetic analyses only recovered some of these character states as synapomorphies of this genus (see diagnosis and Appendix 2). The proposal of Domínguez (1988) and Hubbard & Domínguez (1988) concerning the intermediacy of Priasthenopus gilliesi with respect to Asthenopus curtus and Asthenopodes picteti is in concordance with our results. Priasthenopus gilliesi resulted sister to the Povilla-Asthenopus clade, presenting some plesiomorphic character states shared with Asthenopodes.

Male

Female and eggs of Asthenopus species are strongly similar. They may be identified by comparison with co-occurring males. Nevertheless the eggs extracted from female adults or mature nymphs may be keyed as follows:

Nymphs (only 3 species known, almost undistinguishable, the characters below should be confirmed with the study of more material)

Photographs of the type at the British Museum were studied.

Additional material. Two male imagos (IBN, slide 480) from COLOMBIA, Amazonas, Leticia, caño km 15, S 4°5'41" − W 69°59'1", 93 m, 11.ii.1999, light trap 4−6 h, E. Domínguez, M.C. Zúñiga & C. Molineri cols.; male imaginal slides (FAMU) from BRAZIL, Amazonas, Careiro Island, Divinopolis, SE of Manaus, 29.vii.1961, E.J. Fittkau; 2 male and 1 female pharate subimagos (IBN642CM-eggs, 643-female, 644-male) from BRAZIL, Amazonas, São Paulo de Olivença, Bom Sucesso, 4.ix.2003, (aprox. S 3°28' − W 68°59').

Asthenopus curtus is the type species of the genus, and is known from adults of both sexes, nymphs and eggs. Nine autapomorphies were recovered in the cladistic analysis, and are useful to diagnose the species (see Appendix 2). The following combination of characters is useful to distinguish A. curtus from the other species of the genus: 1) male FW 10.0, female FW 14.0–18.5; 2) male foreleg length 0.69–0.74 times the length of FW; 3) pronotum width/length ratio: 2.0–2.3 (male), 2.7–3.0 (female); 4) 18–25 marginal intercalary veins present on the entire margin of forewings (Fig. 16A–B), 2–3 times shorter than distance between longitudinal veins in male (not anastomosed), hind wings with marginal intercalaries in at least 4 spaces between main veins; 5) male FW with 0 to 1 crossveins between Rs and MA basal to Rs fork; 6) ratio total length/basal width of forceps 5.4 (Fig. 17A–B); 7) penes very sclerotized, contrasting strongly with the remaining genital parts, apex projecting acutely; a deep furrow separates penis lobe from thumb, median remnant of styliger plate subrectangular without marked projections, pedestals subrectangular and large, outer margin projecting posteriorly on outer margin along forceps base; 8) female sternum VIII with anteromedian keel and reduced sockets as in Fig. 18G; 9) egg ratio maximum width of egg/maximum width of PC 1.1–1.3, cap formed by 3–5 filaments, chorionic plates separated by smooth chorion (Fig. 18A); 10) nymphal mandible: ratio total length of mandible/mandibular tusk length 1.6–1.7; 11) space between the subbasal and the submedian tubercles in inner margin of left mandibular tusks is short and concave (Fig. 14A).

Male imago. Length: body, 8.0–8.7; FW, 10.0; HW, 4.4; foreleg, 7.5; cerci, 33.0–35.0. General coloration yellowish light brown. Head whitish, heavily shaded black dorsally, paler on posteromedian zone of occiput, black shading extending anteriorly on frons as two parallel lines surrounding median ocellus. Antennae pale, slightly shaded gray on dorsum. Thorax. Pronotum yellowish translucent completely shaded gray, darker on anterior ring; paler on two transverse lines, one separating anterior and posterior rings and another more posterior and obliquely transverse; pleurae shaded with black, sternum with a median gray macula. Pronotum width/length ratio: 2.0–2.3. Mesonotum whitish yellow (or brownish in some males) with a black median triangle between posteroscutal protuberances, metanotum similar in color, also shaded black posteromedially; mesopleurae and sterna paler, shaded with black along anterior margin of katepisternum. Legs yellowish white shaded with gray dorsally on all coxae, femora and tibiae; foretarsal segment 1 blackish (Fig. 20G), remaining tarsal segments paler shaded with gray distally, claws grayish thin troughout. Wings (Fig. 16A–B). Membrane hyaline shaded very slightly with brownish near anterior margin and turning whitish translucent towards apical zone of C–Sc areas; veins translucent shaded with brown. Abdomen yellowish white shaded extensively with grayish brown dorsally, darkening very slightly towards rear segments. Sterna pale very slightly shaded gray, shaded stronger on mediolongitudinal line near anterior margin of sterna VIII–IX, this line is blurred posteriorly; a grayish black triangular mark is present at each side of this line, on anterior margin of sterna VIII–IX; sternum X shaded black except medially. Genitalia (Fig. 17A–B): median remnant of styliger plate and pedestals yellowish, forceps whitish translucent shaded gray along outer margin, penes dark orange with whitish base. Caudal filaments whitish, shaded gray at base of terminal filament.

Female adult. Length: body, 10.5–13.2; FW, 14.0–18.5; HW, 5.7; cerci, 5.8–7.0. Morphologically very similar to female adults of A. angelae described in detail in de Souza and Molineri (2012). Here only those characters that differ from the cited description are mentioned. Pronotum almost 3 times wider than long, width/length ratio = 2.7–3.0 (see continuous characters in phylogenetic matrix, Appendix 3). Mesonotum uniformely brownish (cuticular pigmentation), almost without gray markings (dermic pigments). Female sternum VIII with anteromedian keel and reduced sockets as in Fig. 18G. Cercus about half the length of FW, cercus length/FW length: 0.4–0.5.

Eggs (Fig. 18A). Length, 200–220 µ; width, 130–155 µ. Two polar caps (maximum width, 110–120 µ), formed by 3–5 very long coiled threads. Chorionic surface smooth with relatively large subcircular chorionic plates, the plates are regularly spaced and some of them are divided in two or three subequal parts.

Mature nymph. Length of male: body, 9.5–9.7; cercus, 7.0; terminal filament, 5.0. Length of female: body, 17.0; cercus, 8.0; terminal filament, 7.0. Only characters that differ from A. angelae are given here, refer to that description for more detailed information. Head (occipital area) dorsally brownish uniformly shaded with gray. Mouthparts. Left mandibular tusks with a relatively shorter space between the large subbasal tubercle and the smaller subdmedian one, this space is somewhat C-shaped (Fig. 14A). Right mandible with distal corner of mola strongly protruding. Thorax. Mesonotum uniformly brownish (cuticular) without strongly gray-shading on carinae (Fig. 4E). Legs and paraprocts identical to those on Fig. 15.

Amazonas River from Leticia (Colombia) to Manaus (Brazil).

Much confusion exists in the literature concerning this species. Many authors mention A. curtus but from missidentified material. For example Ulmer (1942) described and illustrated (as A. curtus) a pair of males of Asthenopus angelae. The material from Ecuador studied by Domínguez (1988) proved to be a different but related species (A. magnus sp. n.). Berner (1978) synonymized A. curtus with A. amazonicus, showing that the differences between both species were only attributable to sexual dimorphism, but he was working with A. angelae males (de Souza and Molineri 2012). Nevertheless, Berner conclusions were correct given that sexual dimorphism in FW venation is present in both species. As it is impossible to assign any specimen to A. amazonicus, we prefer to treat it as synonym of A. curtus, as Berner proposed. Actually, only one specimen from previous works is positively determined as A. curtus: the type, studied by Eaton (1883) and illustrated by Kimmins (1966). We add here some other records from the Amazonas River: a pair of males from Colombia, some reared nymphs from Brazil and Fittkau’s slides at FAMU. These male imagos show the characteristic genitalia of the holotype of A. curtus, with extremely wide forceps, long penis lobes and slender and very acute apical spines, and the more or less uniform brownish mesothoracic coloration (an exception of this last character are the males from Colombia-Leticia, much paler). The egg of A. curtus (Fig. 18A) is similar to that of A. hubbardi (Fig. 18D), in the shape and relative large size of the disk-like structures that leaves exposed only a reduced surface of smooth chorion. On the contrary the egg of A. angelae presents smaller disk-like structures with a larger surface of smooth chorion among them (Fig. 18B).

As the result of the present study, the female adult, egg, and nymphal stages are described here for the first time. Previous descriptions of female and nymphs in the literature were done from specimens of A. angelae or other species but are not useful to clearly distinguish the species.

Holotype (IBN) male imago from Ecuador, Napo Province, Laguna Limon Cocha, 250 m, 6.iv.1984, E. Domínguez col.(aprox. S 0°24' − W 76°38'). Paratypes, same data as holotype, separated in 16 vials including: 1 nymph dissected (parts in alcohol), 2 male imagos (parts on slides: IBN-2–64ED, IBN-2–70ED), 1 male imago (IBN-2–67ED), 2 nymphs/3 exuviae/1 pharate male, 1 pharate male subimago and nymphal cuticle (IBN483CM), 1 male imago (IBN481CM), 3 nymphal exuviae (IBN640CM, IBN641CM), 7 nymphal exuviae (1 at FAMU, 1 at CZNC), 5 female adults (used for SEM); 7 male imagos (used for the description; 1 at FAMU, 1 at CZNC), 20 male subimagos, 10 female adults, 1 male and 8 female subimagos, 10 female adults, 9 female adults (1 at FAMU, 1 at CZNC), 1 female adult (IBN-2–71ED). All the material is deposited in IBN except otherwise indicated.

Asthenopus magnus, known from all the stages, can be distinguished from other species in the genus by the following combination of characters (also see the six autapomorphies in Appendix 2): 1) male FW 9.0–10.1, female FW 16.0–17.5; 2) forelegs of male 0.69–0.73 × the length of FW; 3) pronotum width/length ratio: 1.7–2.4 (male), 2.0–2.2 (female); 4) FW (Fig. 16C) with 6–14 relatively short marginal intercalaries (21–28 in female), hind wings with 2–4 (5–7 in female) marginal intercalaries; 5) male FW with 0 to 2 (2–3 in female) cross veins between Rs and MA basal to Rs fork; 6) forceps relatively slender, ratio length/basal width 6.1–6.2 (Fig. 17G); 7) penes tubular and robust, with well developed thumb, curved ventro-medially, with apex projecting medially as a distal spine, furrow separating penis lobe from thumb reduced; median remnant of styliger plate subrectangular, pedestals subrectangular to subovate, relatively large (Fig. 17G); 8) female sternum VIII with reduced, not distinguishable female sockets, but with a long anteromedian keel (Fig. 18F); 9) eggs (Fig. 18E) ratio maximum width of egg/maximum width of PC 1.2–1.3, cap formed by 4–5 filaments, chorionic plates separated by smooth chorion; 10) nymph, ratio total length of mandible/mandibular tusk length 1.4–1.5; 11) inner margin of left mandibular tusk with subbasal and submedian tubercles well separated (Fig. 14C, E).

Male imago. Length (mm): body, 9.0–10.5; FW, 9.0–10.1; HW, 4.0–4.8; leg I, 6.5–7.4; cerci, 37.0. General coloration yellowish light brown. Head whitish shaded black dorsally on pale median mark on hind margin and along inner margin of eyes; frons pale except paired submedian black lines. Antennae: whitish shaded diffusely with gray on scape and apex of pedicel; length (mm): scape 2.25, pedicel 1.5, flagellum 7.25. Thorax. Pronotum yellowish translucent shaded with black dorsally except at pale median membrane between both pronotal rings, on mediolongitudinal line and along margins; the black shading presents many scattered and small pale spots. Pronotum width/length: 1.7–2.4. Meso- and metanotum yellowish shaded with grayish on carinae, posteromedian triangular mark (on mesonotum), and scutellum (both). Thoracic pleurae and sterna paler, shaded gray on pleural sclerites. Legs yellowish white, shaded gray on all coxae. Leg I shaded gray almost completely, stronger on femur and tibia, paler on tarsal segments (Fig. 20J). Legs II–III shaded with black dorsally on apex of femora and entire dorsum of tibiae, rest pale. Wings (Fig. 16C–D). Membrane hyaline shaded grayish near costal margin on basal half, more whitish apically; all veins translucent completely shaded gray; 1–3 cross veins between MA and R, basad to R stem. Abdomen yellowish white shaded with gray and black dorsally except on median and lateral zones and intersegmental membranes. Median pale areas on terga II–IX are oval and are surrounded by darker pigments, a delicate mediolongitudinal black line is present but sometimes is only visible on tergum IX. Sterna whitish turning yellowish laterally and on sternum IX, shaded gray on paraproct and basally to terminal filament. Genitalia (Fig. 17G): forceps whitish, penes yellowish white. Cerci whitish very slightly shaded with gray.

Female imago. Length (mm): body, 15.5–19.5; FW, 16.0–17.5; HW, 5.2–7.8; cerci, 3.5–5.0. Pronotum width/length: 2. Morphologically very similar to A. curtus and A. angelae, the last is described elsewhere (de Souza and Molineri 2012). Color pattern similar to male but more strongly marked, exceptions follows: pale median mark on occiput longer, reaching median ocellus; wing membrane tinged with yellowish near costal margin and base, all veins shaded brown; shading on abdominal terga more extended, shaded widely gray except medially, tergum VIII–IX (sometimes also II–VII) with a black line in the pale median zone; medial margins of gill sclerites on abdominal sterna II-VII and lateral margins of sterna VIII–IX grayish; Sternum VIII with keel as in Fig. 18F. Cerci yellowish, 0.3–0.4 the length of FW.

Eggs (Fig. 18E). Length, 250–285 µ; width, 145–160 µ. Two polar caps (maximum width, 115–135 µ), formed by 4–5 very long coiled threads. Chorionic surface smooth with relatively large subcircular chorionic plates, the plates are regularly spaced and some of them are divided in two or three subequal parts.

Nymphs. Length of male (mm): body, 10.0–11.0 mm; cerci, 7.0–8.0; terminal filament, 5.0–5.5. Length of female (mm): body, 17.0–20.0 mm; cerci, 4.0–5.0; terminal filament, 5.0. General coloration brownish. Head (Fig. 14I–J) yellowish brown extensively shaded with grayish brown, darker on a band between ocelli, occiput with a profuse netted grayish pattern, except on paler median zone and along inner margin of eyes; paler areas also present basally to antennae and around median ocellus. Antennae yellowish white, length (mm): scape (0.5), pedicel (0.28), flagellum (2.0). Mandibular tusks with relatively large space between large basal tubercle and smaller subdistal tubercle, not C-shaped as in A. curtus but in the form of broad “C” or bracket-shaped (Fig. 14C, E, G). Thorax. Anterior ring of pronotum (collar) blackish; posterior ring shaded gray except on mediolonitudinal line and a pair of sublateral pale marks; pronotal membranes whitish. Mesonotum brownish, lighter toward apex of wingpads. Thorax ventrally whitish. Legs (Fig. 15A–D) whitish yellow shaded with gray on coxae, apex of femora and along tibiae; foretarsal claw with 20 (male) to 31 (female) denticles in a marginal row, increasing in size distally; distal region of hind femur with a group of ca. 60 (male) to ca. 100 (female) stout spines. Abdomen. Terga shaded gray dorsally, except on paler median band, a thin black median line is present inside this pale area on tergum IX–X; lateral zones of terga below gills, pale. Gills whitish almost completely shaded with gray, darker on outer (exposed) zones. Sterna whitish shaded with gray on lateral margins of sterna VIII–IX, and with grayish black on paraprocts (Fig. 15E). Caudal filaments yellowish.

From Latin “magnus” meaning “large”, noun in apposition. The name alludes to the general size of the individuals, mainly the female adults.

Only known from the type locality in Napo (Ecuador).

The type series described here as A. magnus were previously treated as A. curtus (Domínguez 1989: 173; Domínguez et al. 2006: 561), and used to record the latter species in Ecuador. As a result of our study we found some characters distinguishing these specimens as a new species, and thus A. curtus is no more considered to be present in that country. Nymphs and adults of both sexes were associated by nymphal exuviae and adults caught at the moment of emergence.

holotype male imago (slide IBN479CM) from Colombia, Amazonas, Puerto Nariño, Loreto Yacu, S 3°44'26" − W 70°27'19", 5.ii.1999, luz 6−8 h, M.C. Zúñiga, E. Domínguez and C. Molineri cols.; and paratypes: 1 female imago (slide IBN574CM) same data as holotype; and 1 male imago (slide IBN605CM) from Colombia, Amazonas, Puerto Nariño, Lago Tarapoto, S 3°47'47" − W 70°25'17", 4.ii.1999, light trap 18–20 hs, M.C. Zúñiga, E. Domínguez and C. Molineri cols. Holotype deposited in MUSENUV, paratypes in IBN.

Asthenopus hubbardi, known from adults of both sexes, can be distinguished from the other species in the genus by the following combination of characters (one autapomorphy is listed in Appendix 2): 1) male FW 7.8–9.2 (Fig. 16E–F), female FW 13.0; 2) forelegs of male 0.57–0.65 × the length of FW, apex of foretibia with stout spines (Fig. 20I); 3) pronotum width/length ratio: 2.1 (male), 2.7 (female); 4) FW with 4–14 relatively short marginal intercalaries (ca. 20 in female), hind wings without marginal intercalaries (present in all spaces in female); 5) FW with 2–3 cross veins between Rs and MA basal to Rs fork, in both sexes; 6) forceps relatively slender, ratio length/basal width 4.7–6.0 (Fig. 17C); 7) penes tubular and robust, furrow separating penis lobe from thumb well marked, median remnant of styliger plate subrectangular, pedestals subrectangular to subovate, relatively large (Fig. 17C); 8) female sternum VIII with reduced female sockets, anteromedian keel present (Fig. 18H); 9) eggs ratio maximum width of egg/maximum width of PC 1.1–1.3, cap formed by 3–8 filaments, chorionic plates almost contiguous (Fig. 18D).

Male imago. Length (mm): body, 7.0–7.1; FW, 7.8–9.2; HW, 3.3–4.0; leg I, 5.1–5.2; cerci, 22.0–23.0. General color whitish brown. Head whitish shaded black dorsally except thin line along hind margin, with a pair of blackish short lines anteriorly to median ocellus. Antennae whitish shaded gray at margins of scape. Thorax. Pronotum ratio width/length: 2.1. Pronotum yellowish translucent widely shaded black, except on pale transversal line between anterior and posterior rings, and on mediolongitudinal line of posterior ring, and posterolateral oblique dashes. Meso- and metanotum whitish yellow shaded black on carinae and margins, also shaded on black on posteromedian triangular zone. Legs whitish shaded gray on dorsum of foreleg, all coxae, and on legs II–III on apex of tibiae and dorsum of tarsi; foretarsal segment 1 is shown in Fig. 20I. Wings (Fig. 16E–F) membrane hyaline except apically whitish on C and Sc areas, veins translucent yellow turning hyaline distally, except basal 2/3 of veins Sc and R₁ yellowish; 2 cross veins between R and M basally to R stem. Abdomen whitish shaded almost completely with brownish gray except at pale intersegmental membranes, oblique dashes on lateral zones of terga I–VIII, submedian pale spots on anterior margin of terga II–IX, and single posteromedian pale spot on terga III–IX. Genitalia (Fig. 17C) whitish except penes yellowish. Caudal filaments whitish, slightly shaded gray on basal segments of cerci.

Female imago. Length (mm): body, 10.5; FW 13.0; HW, 5.0, cerci 3.8. Similar to male, shaded dorsally more uniformly and markedly. Pronotum ratio width/length: 2.7. FW with 4 cross veins between R and M basal to R stem (none of them just below fork). Abdominal sternum VIII with keel as in Fig. 18H. Cerci yellowish turning whitish apically; ratio length cercus /FW = 0.3.

Eggs (Fig. 18D). Length, 210–245 µ; width, 135–150 µ. Polar caps (maximum width, 110–135 µ) formed by 3–8 long coiled threads. The chorionic plates are almost contiguous, leaving a reduced smooth chorionic surface among them (Fig. 18D).

The species is named for Mike Hubbard who has contributed significantly to the understanding of mayflies throughout the world.

Two near localities in the Amazonas River from Colombia.

This species is very similar to A. angelae, and both were collected in the same lightraps in Colombia, nevertheless they can be separated because A. hubbardi shows translucent veins in the wings (brownish in A. angelae); in most specimens a cross vein is present just below R fork in A. angelae (more basal or distal in A. hubbardi), and the penes are shorter and well separated from the basal thumb in A. hubbardi (similar to A. curtus). Asthenopus hubbardi is further characterized because foretibia (Fig. 20I) presents strong marginal spines (weak or absent in A. curtus).

Holotype male imago (slide IBN473CM) from Argentina, Corrientes, Parque Nacional Mburucuya, Selva Misionera (sector 6), luz, 29.iii.2001, F. Navarro col.

Additional, non-type material. One reared female subimago (IBN524CM) and nymphal cuticle (IBN639CM) from Argentina, Corrientes, Laguna Brava, 27.i.1977, Poi de Neiff col. (egg in Fig. 18C extracted from this female); and 1 male imago (IBN638CM) from Uruguay, Salto, near Salto Grande, frente a Isla del Paredón, 20−21.I.1975, luz 22hs; 4 male and 10 female imagos from Brazil, São Paulo, Luiz Antonio, 10.iv.1991, C.G. Froelich col. (MZSP); 1 male and 4 female imagos from Brazil, São Paulo, Luiz Antonio, Reserva Jatai, 9.iv.1990, C.G. Froelich col. (MZSP).

Asthenopus guarani, known from all stages, can be distinguished from the other species in the genus by the following combination of characters (seven autapomorphies are detailed in Appendix 2): 1) male FW 8.0–9.0 mm (Fig. 16G–H), female FW 16.0 mm; 2) Ratio FW/foreleg length 1.4–1.8; 3) pronotum width/length ratio: 2.15 (male), 3.1 (female); 4) male FW with 4–6 marginal intercalaries (24–26 in female), slightly shorter than the separation of main veins, HW (Fig. 16I) with 2–4 marginal intercalaries (8–9 in female); 5) male FW with 0–2 cross veins between Rs and MA basal to Rs fork (3 in female); 6) forceps relatively slender, ratio length/basal width 4.8–5.8 (Fig. 17E–F); 7) penes tubular and slender, furrow separating penis lobe from thumb well marked; median remnant of styliger plate with lateral rounded lobes as in Fig. 17E). 8) female sternum VIII with reduced, not distinguishable female sockets, but with a long anteromedian keel; 9) eggs (Fig. 18C) ratio maximum width of egg/maximum width of PC 1.2, cap formed by 3–8 filaments, disk-like structures well separated by smooth chorion, with 2–3 small disks beneath each larger disk (Fig. 18C); 10) nymph, ratio total length of mandible/mandibular tusk length 1.4; 11) inner margin of left mandibular tusk with subbasal and submedian tubercles well separated (similar to Fig. 14C, E).

Male imago. Length (mm): body, 6.5–9.0; FW, 8.0–9.0; HW, 3.6–4.0; foreleg, 5.1–5.4; cerci, 25.4–28.0. General coloration yellowish white. Head whitish shaded with black dorsally except on hind margin and posteromedian pale mark; frons pale shaded with a pair of black submedian longitudinal lines; venter of head pale. Antennae whitish shaded with gray on scape; length (mm): scape 1.75, pedicel 1.25, flagellum 7.5. Thorax. Pronotum whitish translucent, shaded black on anterior ring and lateral margins; posterior rings shaded gray, with darker mediolongitudinal line. Meso- and metanotum yellowish white, shaded gray on scutellum; sterna pale shaded gray only on mesokatepisternum. Legs whitish shaded gray on coxae. Foreleg completely shaded gray, paler on base of tarsal segments 2–5 (Fig. 20H). Middle and hind legs shaded gray on apical half of femora and apical 2/3 of tibiae, tarsi and claws translucent. Wings (Fig. 16G–I). Membrane of both wings hyaline, except shaded with light gray at base of costal margin in both wings and whitish on the apex of costal margin of FW; veins translucent, except Sc and R₁ whitish, all veins shaded slightly gray but more diffusely toward apex. Abdomen whitish, terga uniformly shaded gray, paler laterally and anteriorly to each tergum; sterna whitish. Genitalia (Fig. 17E–F) whitish, penes yellowish white. Cerci whitish translucent.

Female subimago. Length (mm): body, 12.5; FW, 16.0; HW, 6.5; cerci broken off and lost. General coloration orangish yellow shaded widely black. Head dorsally blackish except medial line on occiput, and anteriorly to median ocellus. Thorax. Pronotum width 2.5 mm, total length 0.8 mm; cream shaded black on thin anterior ring, with gray on posterior ring, membranes whitish. Mesothorax orangish yellow with gray markings. Wings translucent whitish, veins whitish except basal half of C, Sc and R yellowish. Abdomen uniformely shaded gray dorsally, except on pale medial line. Sternum VIII with long and thin anteromedian keel. Base of caudal filaments whitish (rest broken off and lost).

Eggs (Figs 18C). Length, 160–200 µ; width, 100–160 µ. Polar caps (max. width 130 µ) formed by 3–8 long coiled threads. The disk-like structures are circular and entire (not partitioned as other species), very well separated by smooth chorion, and when removed, a group of 2–3 very small disks are visible beneath (Fig. 18C).

Nymph (cuticle from reared female described above). Length (mm): body, 16.5 mm; cerci and terminal filament, 7.0 (both broken at apex). Antennae broken off and lost. Mouthparts. Mandibular tusks with relatively large space between the large basal tubercle and smaller subdistal tubercle, not C-shaped (similar to Fig. 14C, E, G). Legs. Foretarsal claw with a row of 26–29 denticles. Apex of hind femur with a group of 90–100 stout acute spines.

The name refers to one of the etnic groups inhabiting the area where the specimens were collected.

Argentina (Corrientes), Brazil (Sao Paulo), Uruguay (Salto).

This species is very distinctive, not only by the long and slender penis lobe, but also because of the presence of relatively long marginal intercalary veins (in male FW and HW). The reared female subimago is tentatively associated with the male, because of similarity in coloration and shared distributional range. Eggs extracted from this female show also some differences from the other known in the genus, mainly the larger extent of smooth chorion around the plates and the presence of small disks below the larger ones (Fig. 18C).

(see de Souza and Molineri 2012). Additional records: 13 nymphs (including 3 male and 1 female pharate subimago) from BRAZIL, Amazonas, Codajás, Urucurizinho, Lago Cuxuará (A10-lago Aquapi 14), 15.ix.2003 (slides IBN645CM and IBN646CM) (aprox. S 3°55' − W 62°3'); and 1 pharate male subimago from BRAZIL, Amazonas, São Paulo de Olivença, Bom Sucesso (A02-agapito), 9.iv.2003 (slide IBN606CM) (aprox. S 3°28' − W 68°59'). Deposited in INPA (4 nymphs), CZNC (6 nymphs), IBN (3 nymphs).

Only one autapomorphy was recovered in our analysis for A. angelae, a change in the ratio A (total length forceps)/E(basal width) from 6.2 to 6.5–7.1 (i.e., forceps become slightly slender). This species can be recognized by the following combination of characters: 1) FW size male 7.0–10.0 mm (Fig. 16J–K), female 12.0–17.0 mm; 2) ratio FW/foreleg length 1.4–1.6; 3) pronotum width/length ratio: 2.0–2.2 (male), 2.2–2.9 (female); 4) 5–11 imv present in male FW, relatively short and poorly anastomosed; 5) male FW with 1–4 cross veins between Rs and MA basal to Rs fork; 6) ratio total length/basal width of forceps 6.5–7.1 (Fig. 17D); 7) penes tubular, with well developed thumb, curved ventro-medially, with apex projecting acutely, furrow separating penial lobe from thumb reduced or absent (Fig. 17D), pedestals subrectangular to subovate, relatively large; 8) female sternum VIII with anteromedian keel and reduced sockets similar to other species; 9) eggs (Fig. 18B) with 3–5 threads on polar caps, ratio maximum width of egg/maximum width of PC 1.1–1.3, chorion with smooth area around rounded disks; 10) nymph, ratio total length of mandible / mandibular tusk length 1.6; 11) space between the subbasal and the submedian tubercles relatively long and straight (Fig. 14B).

Argentina, Bolivia, Brazil, Colombia and Peru.

This species was recently described from all the stages (de Souza and Molineri 2012), and in our analysis it appears as sister to A. magnus, but with relatively low support. In the original description, nymphs and female adults were not distinguished from A. curtus, because of lack of characters. We proposed (above in diagnosis) some characters that should be checked and confirmed with the study of more material. We suggest that specific identification of this and the other species of the genus should be confirmed when possible with the study of male genitalia. Peru was mentioned by de Souza and Molineri (2012) in the list of material but not in the distribution, so here it is added in that section.

Figure 21. 

Optimization of selected continuous characters. A Char. 1 (1.416 steps) Ratio length second foretarsal segment/foretibia B Char. 27 (3.791 steps) Nymph, width of tusk as ratio between length of tusk (T) / width of tusk at the base (W) C Char. 13 (23.811 steps) Ratio A(total length forceps)/E(basal width). Arrows indicate increments or decrease in the characters (up and bottom directed arrows, respectively); red arrows indicate a marked change for the node.

Figure 22. 

Optimization of selected continuous characters. A Char. 0 (3.873 steps) Ratio subapical width of foretibia/subbasal width of tarsal segment 2 B Char. 5 (2.344 steps) Ratio length FW/HW C Char. 2 (4.148 steps) Ratio FW/foreleg length D Char. 18 (2.225 steps) female prothorax, width/length. Arrows indicate increments or decrease in the characters (up and bottom directed arrows, respectively); red arrows indicate a marked change for the node.

Figure 23. 

World map indicating the distribution of the three subfamilies in Polymitarcyidae. Additionally, barriers 1 (separating the genus Asthenopus vs Povilla + Languidipes) and 2 (separating Povilla from Languidipes) found in the biogeographical analysis are marked.

Figure 24. 

Map of South America (in part) showing the distribution of Asthenopus species. Barrier 3 found in the biogeographical analysis separates A. guarani from the remaining species in the genus. Barrier 4 (see detail at left bottom) separates A. hubbardi from the rest; and Barrier 5 separates A. magnus from its sister A. angelae.

Figure 25. 

Map of South America (in part) showing the distribution of Asthenopodes species. Barrier 6 found in the biogeographical analysis separates A. chumuco from the remaining species in the genus.